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Contributions to Zoology, 76 (2) 63-85 (2007)

Revision of the genus DiaphoroceraHeyden, 1863 (Coleoptera, Meloidae, Cerocomini)

Federica Turco and Marco A. BolognaDipartimento di Biologia, Universit Roma Tre, viale Marconi 446-00146 Roma, Italy, [email protected], bologna@

uniroma3.it

Key words: Coleoptera, Meloidae, Diaphorocera, taxonomy, phylogeny, morphology, faunistics, biogeography,Palaearctic region.

Abstract

Diaphorocera, a Saharo-Sindian genus belonging to the tribeCerocomini, is revised and a new synonymy is proposed. A cla-distic classification is proposed as well, on a set of adult morpho-logical characters. The available bionomical records, both origi-nal and from literature, concerning phenology, elevation, habitatpreference, and host plants, are summarised. Adult morphologyof all species is described and figured, the catalogue of localitieswith maps of distribution is reported, and a biogeographicalanalysis is proposed.

Contents

Introduction ...................................................................................... 63Material and methods ..................................................................... 64Results ............................................................................................... 65 Bionomics ................................................................................... 65 Classification ............................................................................. 65Key to the species ofDiaphorocera ........................................... 67Taxonomy ......................................................................................... 68 GenusDiaphorocera................................................................ 68 Group obscuritarsis.................................................................. 69 Diaphorocera carinicollis................................................. 69 Diaphorocera johnsoni...................................................... 70

Diaphorocera obscuritarsis ............................................. 71 Group hemprichi ....................................................................... 73 Diaphorocera chrysoprasis .............................................. 73 Diaphorocera hemprichi................................................... 75 Diaphorocera promelaena................................................ 76 Diaphorocera sicardi......................................................... 78 Diaphorocera peyerimhoffi............................................... 78Biogeography ................................................................................... 80Acknowledgements ........................................................................ 83References ........................................................................................ 83Appendix 1 ....................................................................................... 85Appendix 2 ....................................................................................... 85Appendix 3 ....................................................................................... 85

Introduction

The family Meloidae (Coleoptera, Tenebrionoidea) wasrecently revised cladistically (Bologna and Pinto, 2001)and four subfamilies were recognised: Eleticinae,Meloinae, Tetraonycinae and Nemognathinae (Pintoand Bologna, 1999; Bologna and Pinto, 2002). Blisterbeetles are phytophagous, feeding usually on leavesand/or flowers of several plant families, and are char-acterised by cantharidin production and hypermeta-morphic development, except in the primitive sub-family Eleticinae (Pinto et al.,1996; Bologna et al.,

2001). Larvae feed on the provisions and larvae ofHymenoptera Apoidea, or on grasshopper (Acridoidea)eggs (Bologna, 1991). Within the subfamily Meloinae, our interest wasrecently addressed to the Old World tribe Cerocomini,which is composed of five genera (Bologna and Pinto,2002): Cerocoma Geoffroy, 1762 (about 27 species),DiaphoroceraHeyden, 1863 (8 spp.),AnisarthroceraSemenov, 1895 (2 spp.),RhampholyssaKraatz, 1863(2 spp.) and Rhampholyssodes Kaszab, 1983 (1 sp.).Bologna and Pinto (2001) used larval and adult char-acters to reconstruct the phylogeny of Meloidae, but

the placement of the tribe Cerocomini, then repre-sented only by Cerocoma schreberi, remained unre-solved. The authors recognised two larval synapomor-phies for the tribe, then confirmed for other species ofCerocoma(Di Giulio et al., 2002) and also forDiapho-rocera chrysoprasis(Turco et al., 2006). Moreover, themonophyly of the tribe is strongly supported by adultsynapomorphies, as the position of antennae (socketsdistant from eyes, placed below or on the frontal su-ture), the epigamic modifications of male head, anten-nae, maxillary palpi and fore legs, the labrum elongate


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64 F. Turco and M.A. Bologna Taxonomy and biogeography of Diaphorocera (meloid beetles)

and longitudinally furrowed or carinate, the shape ofmouthparts and the endophallic structure (Bologna,1991; Turco et al., 2003). The biology of the tribe is poorly known with theexception of some notes on host plants (Bologna, 1991),

on sexual behaviour of the genus Cerocoma(Turco etal., 2003), and on larval hosts of few Cerocomaspecies(see Bologna, 1991 for a synthesis). Pre-imaginal stagesof Cerocomini are known only for six species of Cero-coma (Di Giulio et al., 2002 for a review; Turco andBologna, unpublished), and oneDiaphorocera(Turcoet al., 2006). Eggs are laid in holes dug by the femalein the ground as in other Meloinae, and the first instarlarvae (triungulins) are not phoretic, actively reachingthe hosts nest where they develop to adult stage. As faras is known, triungulins feed on larvae and honey or oneggs and food stored by Hymenoptera Aculeata (Bolog-

na, 1991 for a review). After the first studies on the larval morphology (DiGiulio et al., 2002; Turco et al., 2006), our project wasaddressed to the revision of the tribe Cerocomini as awhole. The present paper is focused on the study of thegenusDiaphorocera; another contribution is addressedto the classification of the five genera and to the revisionofAnisarthrocera,RhampholyssaandRhampholyssodes(Turco and Bologna, unpublished); the third study willbe devoted to the revision of the more speciose genusCerocoma(Turco and Bologna, unpublished). The genusDiaphorocerais a typical Saharo-Sindianelement, distributed in the whole Northern Africa,southern Middle East (Palestine and Israel), ArabianPeninsula, and in southern Iran. This genus was neverrevised: Bedel (1895) published the first key to the spe-cies, updated by Kaszab (1951; taken up by Dvorak,1989), after which one new species and one new subspe-cies were described (Kocher, 1954; Kaszab, 1983).

Material and methods

During our field researches we obtained ecologicalinformation on host plants and phenology on threespecies (D. chrysoprasis, D. hemprichiof both subspe-

cies, D. johnsoni). We reared these species and weobtained the triungulin ofD. chrysoprasis, describedin a separate paper (Turco et al., 2006). For the taxonomical study we examined 396 speci-mens representing all theDiaphorocera species, withat least one male of each species, including typicalmaterial, as reported in Table 1. The examined speci-mens are preserved in the following collections (as-sociated acronyms reported in the text): CB = M. A.Bologna coll., Universit Roma Tre, Roma; CBA =J. Batelka coll., Prague; CK = S. Krejcik coll., Unicov,Czech Republic; CP = J. D. Pinto coll., University of

California, Riverside; CR = J. C. Ringenbach coll.,Paris; CS = D. Sechi coll., Cagliari; ISR = InstitutScintifique, Rabat; UCD = University of CaliforniaMuseum, Davis; MCSN = Museo Civico di StoriaNaturale G. Doria, Genova; MHNG = MusumdHistoire Naturelle, Genve; MNHN = Musum Na-tional dHistoire Naturelle, Paris; MVE = Museo diStoria Naturale, Venezia; HNHM = Hungarian NaturalHistory Museum, Budapest; OUM = Oxford Univer-sity Museum, Oxford. For the phylogenetical analysis of the eight speciesofDiaphorocera we used the software PAUP 4.0 (Swof-ford, 2002) and 24 adult characters available for allspecies, selected from a larger set previously identified;character 24 was not observed in D. peyerimhoffibe-cause the dissection of types was not possible. Thecharacter matrix with the related list are given in Ap-pendices 1 and 2. According to Bologna and Pinto(2002), the single taxon constrained as outgroup is

Anisarthrocera semirufa (Fair-maire, 1882) (see Classification).Autapomorphic characters were uti-lized for the diagnoses and key only.

As expected, characters displayedsome homoplasies, as indicated byrelatively low consistency index (CI)(Appendix 3). Both binary and multi-state characters were employed, allelaborated as unordered and equallyweighted, and processed by thebranch-and-bound algorithm usingparsimony as optimality criterionand the furthest addition sequenceoption. The accelerated transforma-

Table 1. Types and other specimens examined.Species Types and Other

location specimensD. carinicollis lectotype, 5 paralectotypes MNHN 1D. chrysoprasis 90

D. hemprichi hemprichi 53

D. hemprichi saudita 4 paratypes HNHM 47

D. johnsoni 3 paratypes HNHM 8D. kerimii(syn. ofD. chrysoprasis) holotype MCSND. obscuritarsis holotype, 12 paratypes MNHN, HNHM 76D. obscuritarsisvar. ruficornis syntype MNHND. peyerimhoffi holotype, 'allotype', 1 'cotype' ISR 17D. promelaena 9 paratypes HNHM 73D. sicardi holotype MNHN 4


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65Contributions to Zoology, 76 (2) 2007

tion algorithm (ACCTRAN) was used to optimizecharacters on cladograms. Confidence levels were as-sessed using Bootstrap Analysis and results werecompared by generating a strict consensus cladog-ram.

Results

Bionomics

All the species live in eremic habitats, both on dunes orstony desert (Fig. 1). Species of this genus are strictlydiurnal, and adult occurrence is primarily restricted tospring, as summarised in Table 2. All the species seemto be distributed at low elevation, from sea level to about200 m a.s.l., with a single record ofD. obscuritarsisat

1200 m a.s.l. in southern Morocco.Adults feed on pollen of several plant families; all

personal and literature records are summarised in Table2. Thanatosis was observed in the field inD. hemprichi,D. johnsoniandD. chrysoprasis. In this genus sexualdimorphism is evident mostly on head, antennae, and

sometimes also on fore tibiae, but courtship behaviourhas never been studied, and we only quickly observedon flowers the dorsal and the subsequent linear phasesofD. chrysoprasiscopulation. As previously indicated,the larval hosts and the pre-imaginal development are

unknown, except the triungulin of D. chrysoprasis(Turco et al., 2006).

Classification

This genus was described by Heyden (1863) to includeD. hemprichifrom Egypt, but in the same year a secondspecies was described by Fairmaire (1863) from theopposite side of Sahara, and Bedel (1895) evidenceddefinitively the Saharan distribution of this genus. Sub-sequently other representatives were found also in theArabian peninsula and South Iran, enlarging its range,

which is now similar to that of other eremic genera ofanimals and plants. The phylogenetic relationships of Diaphorocerawithin Cerocomini were never defined and are the objectof a separate contribution (Turco and Bologna, unpub-lished). Larval features of Cerocomini are until now

Fig. 1. Habitat ofD. chrysoprasis, Tunisia, 10 km SW of Tozeur on road P3.


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uninformative, because of their scarcity, homogeneity,and also because pertinent to only two genera.

Kaszab (1951, 1969, 1983) divided the genera onthe base of the number of antennomeres, consideringcorrectly the reduction as a derived condition, but hedid not evaluate other derived features as those of ae-deagus, fore tibiae, etc. The basic number of anten-nomeres of Meloidae is 11, but the reduction is commonparticularly in several genera of the tribe Mylabrini.Two genera of Cerocomini have 11 antennomeres, asa primitive condition: Diaphorocera and Anisar-throcera. Two other genera, namely Cerocoma andRhampholyssodes, have 9-segmented antennae, and afi

fth genus, Rhampholyssa , has 8-segmented anten-nae. Bologna and Pinto (2002) supposed the separation ofAnisarthrocerafromDiaphoroceracould be doubtful.Actually,Diaphorocerais distinct, but its relationshipsmay be looked for examining theAnisarthrocera-com-plex. In fact, this last genus, considered temporarily asmonophyletic by Bologna and Pinto (2002), is actuallypolyphyletic (Turco and Bologna, unpusblished). It in-cludesA. batesiMarseul, 1872, a polytypic species fromIran and Arabia, which is greatly different fromA. sem-irufa (Fairmaire, 1882) from N Somalia and a new

Kenyan species, which probably belong to a new genus(Turco and Bologna, unpublished). These last two spe-cies have unmodified male fore tibiae, unlikeA. batesi,and are slightly similar toDiaphorocera (see also Bo-logna, 1991), from which they differs at least for thesingle hook on both aedeagus and endophallus, in addi-tion to the body colouration. Contrarily to Kaszab (1951),frontal calli are found, more or less developed, in bothAnisarthrocera- complex andDiaphoroceraand cannotbe used to separate them. Waiting for the phylogenetic revision of the tribe

(Turco and Bologna, unpublished), we consider Dia-phoroceraas a primitive genus of the tribe, together

with Anisarthrocera s.str. and the new East Africangenus. Phenetically, this new genus seems more closetoDiaphorocerathan toAnisarthrocera. Relationships amongDiaphoroceraspecies derivedfrom the cladistic analysis, are summarised in a clado-gram (Fig. 2), emerging from a matrix of 8 species (andthe outgroup Anisarthrocerasemirufa) x 24 characters(Appendixes 1 and 2). This most parsimonious clado-gram evidences three main lineages: one including 3species, and the second including a single species, thelast 4 species, subdivided in two minor lineages (see

Table 3 for a list of synapomorphies). The main line-ages represent three monophyletic groups of speciesrecognised primarily by the shape of male fore tibiaeand last antennomere:a The obscuritarsisgroup: D. carinicollis, D. john-

soni,D. obscuritarsis, all with last male antennomeresubquadrate and fore tibiae not modified. The firstcharacter could be considered an apotypic condition,because blister beetles and in particular other Cero-comini, have the last antennomere elongate. Thesecond character is plesiotypic. Relationships amongthe species were not resolved by the cladistic (Fig.

2), but the bootstrap analysis distinguished one cladeincludingD. obscuritarsisandD. carinicollisby avery low confidence value (24).

b Thepromelaenagroup: D. promelaenaonly, char-acterised by the last antennomere elongate as in thehemprichigroup, but distinct because of a differentderived condition of the male fore tibia, modifiedonly distally, and by the integument partially black.The phylogenetic placement of this group seems tobe close to the obscuritarsis group, even if sup-ported by a relatively high consistency index (69).

Table 2.Ecological information on phenology and host plants of adults. Phenology in brackets indicates eccentric records. Ap = Apiace-ae; As = Asteraceae; B = Brassicaceae; F = Fabaceae; R = Resedaceae; T = Tamaricaceae; Z = Zygophyllaceae.Species Phenology Host plantsD. carinicollis late April - MayD. chrysoprasis April - May (February) Chrysanthemum coronarium (As)

D. hemprichi March - May (July, November) Senecio glaucus, Asteriscus graveolens, Anvillea radiata (As);Eremobiumaegyptiacum (B);Reseda arabica (R);Tribulus omanense, Zygophyllummandavillei (Z)

D. johnsoni March - May Zygophyllum mandavillei(Z)D. obscuritarsis April - May (February) Eruca aurea, Farsetia linearis (B);Amni visnaga (Ap);Reseda lutea(R);

Tamarix ramossissima (T)D. peyerimhoffi middle JuneD. promelaena April - early June Retamasp. (F)D. sicardi April - May


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c The hemprichigroup:D. chrysoprasis, D. hemprichi;D. sicardi, D. peyerimhoffi. All these species havethe last antennomere elongate and fore tibiae vari-ously modified (two different apotypic conditions).Relationships among the species appear clearly de-fined. In particular, thefirst two species have the foretibiae greatly modified with a laminar expansion onthe external side, extended to the whole length. Onthe other hand,D. sicardi andD. peyerimhoffirep-

resent a very distinct West Saharan lineage, ex-

tremely supported by the cladistic analysis, charac-terised by some synapomorphies, as the modifiedfrontal calli, the labrum dorsal structure, and thepeculiar shape of male fore tibiae, only bulged in thebasal third.

Key to the species of Diaphorocera

Male1 Foretibiae simple. Antennomere XI subquadrate .. 2- Foretibiae variously modified. Antennomere XI

elongate ..................................................................................... 42 Two black and shiny lines on antennomere XI and

one on antennomere X ...................... D. obscuritarsis- Antennomeres X-XI without lines .............................. 33 Antennomere VII distinctly wider than VI and

slightly narrower than VIII; antennomere I dark.Pronotum slender, anterior portion distinctly nar-rower than temples; anterior grooves only weakly

Fig. 2.Strict consensus tree of two cladograms using matrix in Appendix 1 (see Material and methods).

Table 3.List of synapomorphic characters defining monophylet-ic groups (see Appendix 2 for the definition of characters).(hemprichi-chrysoprasis- 1, 6, 8, 9, 14 sicardi-peyerimhoffi)(hemprichi-chrysoprasis) 2, 20(sicardi-peyerimhoffi) 13, 15, 16, 17, 20(obscuritarsis-carinicollis-johnsoni) 4, 7, 21


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developed. External margin of elytra only slightlysinuate .................................................................. D. johnsoni

- Antennomere VII only slightly wider than VI and aswide as VIII; antennomere I yellow. Pronotum robust,anterior portion only slightly narrower than temples;

anterior grooves deep. External margin of elytraposteriorly greatly sinuate ................... D. carinicollis

4 Head, pronotum, abdomen and antennomere I black........................................................................... D. promelaena

- Head, pronotum, abdomen and antennomere I notblack ........................................................................................... 5

5 Frontal calli with a dorsal keel anteriorly protrudedand pointed; fore tibiae with a basal inflated expan-sion, without laminar expansions ................................ 6

- Frontal calli neither keeled nor anteriorly protruding;fore tibiae with a laminar expansion, at least on theexternal side, extended to the entire length of thetibia .............................................................................................. 7

6 Antennomeres VIII-XI with black lines and spots,anterior portion of X wide, that of VII obliquelytruncate, that of IV deeply incised ........... D. sicardi

- Antennomeres VIII-XI without black lines and spots,anterior portion of X short and thin, that bilobate,that of IV largely incised ................... D. peyerimhoffi

7 Antennomere X about as wide as the length of XI,anterior portion slender and pointed at apex; IX aboutas wide as VIII ............................................. D. hemprichi

- Antennomere X distinctly narrower than the length

of XI. Anterior portion of antennomere IX wide andapically truncate; IX narrower than VIII ..................... .........................................................................D. chrysoprasis

Female

1 Antennomere XI subquadrate ........................................ 2- Antennomere XI elongate ............................................... 42 Fore and middle tarsomeres IV-V dark, basal seg-

ments yellow; trochanters only slightly darkling .... ............................................................................D. carinicollis

- Tarsomeres and trochanters dark, or fore tarsomereI light at base .......................................................................... 3

3 Labrum completely dark; temples slightly divergingposteriad, maximum width of head on temples. Fe-murs and tibiae orange-red ............. D. obscuritarsis

- Labrum dark with the anterior margin orange; tem-ples parallel, maximum width of head on eyes. Fe-murs and tibiae yellow ................................ D. johnsoni

4 Head and pronotum black .................. D. promelaena- Head and pronotum metallic .......................................... 55 Coxae and trochanters black ........... D. peyerimhoffi- Coxae metallic, green or bluish, trochanters yellow

........................................................................................................ 6

6 Body blue, but fore coxae yellow ............ D. sicardi- Body and fore coxae green metallic ........................... 77 Temples elongate, about as long as the eye length,

with subparallel sides .......................... D. chrysoprasis- Temples shorter than the eye length, narrowing

evenly posteriad .......................................... D. hemprichi

Taxonomy

GenusDiaphorocera Heyden, 1863

Diaphorocera Heyden, 1863: 126.

Type species.Diaphorocera hemprichiHeyden, 1863:126, by monotypy.

Diagnosis. The genus Diaphorocerais characterised,as other Cerocomini, by a marked sexual dimorphism.Several male characters do not represent genericsynapomorphies but are present in most species of thegenus, as the modifications of fore tibiae. Because ofthe number (11) of antennomeres, the metallic col-ouration of most part of body, the presence of two hookson both aedeagus and endophallus, Diaphorocera iseasily distinguishable from any other cerocominemeloid. 6-17 mm length. Head most commonly metallic,

green or blue, in one species (D. promelaena)black, insome specimens with a red spot on frons. Labrum, man-dibles, and maxillae very elongate, completely or par-tially orange; galeae fringed. Frons of males with twosmooth calli, just behind the antennae, scarcely raised(D. johnsoni)to very distinct and keeled (D. sicardi, D.peyerimhoffi).Antennae orange or with antennomere Imetallic or black; inserted on the fronto-clipeal suture,11-segmented, subclavate in females, more or lessmodified in males: antennomere I swollen or elongate(respectively in the main lineages); II and III cup-like;IV enlarged, anteriorly normal or bilobate; V and VI

more or less enlarged; VII enlarged with or without adorsal spine extension; VIII-X more or less enlarged andvariously shaped; XI elongate or subquadrate; last anten-nomere similarly shaped in both sexes. Pronotum metallic, green, blue or violet, in one spe-cies (D. promelaena) black; usually elongate, with twooblique impressions on the anterior third in males.Mesopleura not or only slightly depressed just behindthe anterior margin, not touching each other; mesoster-num not differentiated anteriorly; metasternum wide.Elytra metallic, green or blue, flattened, in one species


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evidently sinuate on external side; wings normally de-veloped. Coxae and trochanters yellow, black or metal-lic (green, blue or violet); femurs ant tibiae yellow orreddish; male fore tibiae in one lineage simple, in theother lineages with a laminar external expansion, more

or less extended, or only a short infl

ated expansion;tarsi not modified, yellow or reddish with dark at apexor almost completely dark. Abdomen metallic green, blue, violet, or black; pos-terior margin of male last abdominal sternite emargin-ated, rounded in females. Male genitalia with lobes ofparameres elongate in some species; aedeagus with twodistinct hooks, subequal in size and directed backward;endophallus with sclerotised apical portion bidentate,the apical hook smaller, and outwardly directed.

Commented catalogue. For each species the following

information is indicated: the synonymies; the type local-ity and the type material, also for synonyms; a diagnosisand eventually some taxonomic remarks; the distribu-tion, with the list of localities divided, from West to East,by states and districts, and with the acronym of the col-lection where the material is preserved and/or literaturecitations.

Group obscuritarsis

Diaphorocera carinicollisChobaut, 1921

Diaphorocera carinicollisChobaut, 1921: 298.

Type locality. Touggourt (Chobaut, 1921).

Type material. Male lectotype and five paralectotypes, one maleand four females, were examined (MNHN). The lectotype has thefollowing labels: Touggourt/Mai 1898/ Dr. A. Chobaut (white,printed); Diaphorocera/carinicollis/Chobaut (white, hand-written); LECTOTYPE (red, printed); Diaphorocera/carinicol-lis Chobaut, 1921/lectotype/Jan Batelka desig. 2003 (white,printed). Two paralectotypes have the following labels: Touggourt/Mai

1898/ Dr. A. Chobaut (white, printed);(white, handwritten);carinicollis/Chobaut/types (white, handwritten);MUSUM PARIS/1942/Coll. Dr. A. CHOBAUT (white,

printed); PARALECTOTYPE (red, printed); Diaphorocera/carinicollis Chobaut, 1921/paralectotype/Jan Batelka desig.2003 (white, printed); Touggourt/Mai 1898/ Dr. A. Chobaut(white, printed). (white, handwritten); MUSUM PAR-IS/1942/Coll. Dr. A. CHOBAUT (white, printed); PARALEC-TOTYPE (red, printed); Diaphorocera/carinicollisChobaut,1921/paralectotype/Jan Batelka desig. 2003 (White, printed).The last three female paralectotypes have the same labels, asfollows: Touggourt/Mai 1898/ Dr. A. Chobaut (white, printed); (white, handwritten); PARALECTOTYPE (red, printed);

Diaphorocera/carinicollis Chobaut, 1921/paralectotype/JanBatelka desig. 2003 (white, printed). One of the females lacks right middle tarsus.

Diagnosis.Male. (Figs 3a; 4a; 5a; 6a; 7a, h; 8a-c; 12).13-15 mm length. Body blue-violet metallic, but elytra

green metallic, antennomere I dark, II darkling orangeand II-XI orange, mandibles black, clypeus and labrumdark or partially reddish, maxillary palpi reddish, fronsbetween antennal sockets black, antero-lateral portionof head, anterior to eyes, dark, femurs and tibiae orange,tarsi dark but tarsomere I of fore and middle tarsi orangeand basis of II fore tarsomere reddish. A red frontal spotmore or less evident. Long yellow setae on posterior andventral portions of head, pronotum, scutellum, thoracicpleurites and fore coxae. Frontal calli small and only weakly developed, ante-

rior portion not modifi

ed, scarcely visible on lateral view,posterior margin, on dorsal view reaching about the mid-dle of the inner margin of eye. Temples subparallel,maximum width of head on temples. Labrum about 1,5times longer than clypeus. Maxillary palpomere III thanII, distal end of III obtuse, IV slender and parallel, about2 times as long as the width. Anterior side of antennomereIII with setae longer than those on II; anterior margin ofIV not bilobate, only with an elongate protrusion; ante-rior margin VII without appendix; spine-like setae onanterior apex of antennomeres IV-VII; posterior side ofVI-IX only with short setae; antennomere VII only

slighly wider than VI and slightly narrower than VIII; XIsubquadrate, without chitinous and shiny lines. Pronotum wide, robust, anteriorly slightly narrowed,maximum width on fore side slightly narrower thantemples, fore oblique grooves deep and well developed;middle longitudinal smooth line well evident, formingan apparent carena. External margin of elytra posteri-orly greatly sinuate. Fore tibiae simple, subcylindrical; external spur ofhind tibiae widely spatuled. Parameres in ventral view slender and only slightlyangulated at the base of lobes; lobes elongate (ratiolobes length/parameres length: 0.39); aedeagus as inFig. 8c.

Female. Body green-blue metallic but elytra green-cu-preous metallic, antennae, labrum and maxillary palpidark. Antennomere XI subquadrate.

Taxonomic remarks. This species was recently rede-scribed by Batelka (2004), who figured the male geni-talia and antenna, previously unclearly represented byChobaut (1921).


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Distribution (Fig. 12).Algeria. Algeria (Kaszab, 1951;Dvorak, 1989); Touggourt (Chobaut, 1921; Batelka,2004; MNHN). Tunisia. 20 Km Tozeur on road Nefta-Hazoua (CB); S Kebili (Batelka, 2004; CBA).

Diaphorocera johnsoni Kaszab, 1983

Diaphorocera johnsoniKaszab, 1983: 180.

Type locality. Saudi Arabia, Eastern Prov., UdhailiyahCamp (Kaszab, 1983).

Type material. Three examined paratypes (HNHM) have thefollowing labels. One male: SAUDI ARABIA / Eastern Prov. /D. A. Pitcher (white, printed); Mishash Al Udhailiyah / 13/IV/82 / N 96 (white, handwritten); Paratypus 1983 /Dia-phorocera / johnsoni/ Kaszab (white with red frame, handwrit-ten, except Paratypus printed with red ink); Diaphorocera /johnsoni/ Kasz. / Dr. Z. Kaszab det., 1983 (white, handwritten,except the underlined part of the last line, printed). One maleand one female with the same labels, but 29.IV. / 1982, no. 172and indications of the sex; this male has an additional whitelabel with the genitalia and the last three abdominal segmentsglued.

Fig. 3.Male left antenna, dorsal view: (a)D. carinicollis, (b)D. johnsoni, (c)D. obscuritarsis, (d)D. promelaena,(e)D. peyerimhoffi, (f)D. sicardi, (g)D. chrysoprasis, (h)D. hemprichi. Scale bars = 1 mm.

a

hg

fe

dc

b


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Diagnosis.Male. (Figs 3b; 4b; 5b; 6b; 7b, i; 8d-f; 11).10-12 mm length. Head, pronotum and elytra blue-greenmetallic, thoracic sternites and abdomen blue-violetmetallic, frons between antennal sockets metallic, anten-nomere I, antero-lateral portion of head, anterior to eyes

dark, antennomeres II-XI, maxillary palpi, femurs andtibiae orange, labrum, clypeus, antennomere I and last2-3 tarsomeres dark, anterior margin of labrum yellow,external margin of mandibles reddish. Very long whitesetae on posterior and ventral portions of head, prono-tum, scutellum, thoracic pleurites and fore coxae. Frontal calli small and only weakly developed, an-terior portion not modified, not visible on lateral view,posterior margin, on dorsal view reaching about themiddle of the inner margin of eye. Temples parallel,maximum width of head on eyes. Labrum about 1,5times as long as clypeus. Maxillary palpomere III

shorter than II; distal end of III obtuse; IV slender andparallel, about 2 times as long as the width. Anteriorside of antennomere III with setae as long as those onII; anterior margin of IV not bilobate, only with a shortdigitiform protrusion; VI with a dorsal spur; spine-likesetae on anterior apex of antennomeres IV-VII, veryrobust on VII; anterior margin of VII without appendix;posterior side of antennomeres VI-IX only with shortsetae; VII distinctly wider than VI and slightly nar-rower than VIII; XI subquadrate, without chitinous andshiny lines, apically slightly pointed.

Pronotum slender, maximum width on fore side dis-tinctly narrower than temples, fore oblique groovesonly weakly developed. External margin of elytra pos-teriorly only slightly sinuate. Fore tibiae simple, subcylindrical; external spur ofhind tibiae widely spatuled. Parameres in ventral view slender (length/width isalmost 1), with elongate lobes (ratio lobes length/para-meres length: 0.38); aedeagus as in Fig. 8f.

Female.Colouration as in male but antennae, labrumand maxillary palpi dark; fore margin of labrum light;

femurs and tibiae orange-yellow. Temples narrow andparallel, maximum width of head on eyes; antennomereXI subquadrate.

Distribution (Fig. 11). Saudi Arabia. Saudi Arabia(Dvorak, 1989); Eastern Province, Udhailiyah Camp(Kaszab, 1983; HNHM); Mishash, Al Udhailiyah(Kaszab, 1983); Jaww Dukha (Kaszab, 1983); UnitedArab Emirates. Dubai Emirate, Nahabi (UCD); AbuDhabi Emirate, Al An, Al Jahar-Al Saad on road AlAn-Abu Dhabi (CB).

Diaphorocera obscuritarsisFairmaire, 1885

Diaphorocera obscuritarsisFairmaire, 1885: 38.Diaphorocera obscuritarsisvar. ruficornis Pic, 1923:10 syn. n.

Type locality. Biskra (Fairmaire, 1885).D. obscuri-tarsisvar. ruficornis: Egypt: Massara (Pic, 1923)

Type material. Male holotype (MNHN) and twelve paratypes(HNHM) examined. The holotype has the following labels:Biskra (white, handwritten); obscuri / tarsis (white, handwrit-ten); Diaphorocera/ obscuritarsis (white, handwritten). All theparatypes have the same following labels: Algrie / Biskra / J.Merkl / Mai 1884 (white with thin black frame, printed); coll.R. Oberthr / ex coll Deyrolle (white, printed); Paratypus 1885/Diaphorocera / obscuritarsis/ Fairmaire (white with red frame,handwritten, except Paratypus printed with red ink). The secondparatypes of the line, one female, has additional labels, below theprevious three: Diaphorocera / obscuritarsis (white, handwrit-ten); Fairm type (white, handwritten, upturned). Four females paratypes lack some body parts: both antennae(except the first two antennomeres, still present); hind right leg(except the coxa, still present); the last two tarsomeres of foreright legs (in two females). The female syntype of D. obscuritarsis var. ruficornis(MNHN) was examined. According to Pic (1923), the malesyntype is preserved in the Alfieris collection (Cairo, Egypt), butit was not examined. The female has the following labels: MAS-SARA / MAI (white, printed); 45 (white, handwritten); ex

Chakour (white, handwritten); D. Hemprichi / var (white,handwritten); obscuritarsis / var ruficornis / mihi (white,handwritten); Museum Paris / Coll. M. Pic (white, printed). Some body parts of the female syntype are missing: left an-tenna, except the first two antennomeres; left fore leg, exceptcoax and trochanter; right hind leg; right elytron.

Diagnosis.Male. (Figs 3c; 4c; 5c; 6c; 7c, l; 9c; 8g-i;11). 9-15 mm length. Violet-blue metallic, rarely green-ish, except dark trochanters and tarsi, and orange anten-nae, clypeus, labrum, maxillary palpi, galae, femurs andtibiae; antennomere I slightly dark in some specimens;

fons between antennal sockets black; antero-lateralportion of head, anterior to eyes, dark; a red frontal spotusually present. Frontal calli not clearly visible on lateral view, pos-terior margin on dorsal view reaching about the middleof the inner margin of eye, anterior portion not modi-fied. Temples slightly curved, about as wide as eyes.Labrum about 1,5 times as long as clypeus. Maxillarypalpomere III shorter than II; distal end of III obtuse;IV short and sub-rectangular, about 1,5 times as longas the width. Anterior side of antennomere III with few


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setae, distinctly shorter than those on II; anterior mar-gin of IV not bilobate, only with a digitiform protru-sion; anterior margin of VII with a dorsal slender andpointed appendix; spine-like setae on anterior apex ofantennomeres V-VII; posterior side of VI-IX only with

short setae; one black, chitinous and shiny line on foremargin of X and two oblique in the middle of XI; XIsubquadrate. Pronotum anteriorly with two oblique and deepgrooves. Fore tibiae simple, subcylindrical; external

Fig. 4.Male left maxillary palpus, dorsal view: (a)D. carinicollis, (b)D. johnsoni, (c)D. obscuritarsis, (d)D. promelaena, (e)D. pey-erimhoffi, (f)D. sicardi, (g)D. chrysoprasis, (h)D. hemprichi. Scale bars = 1 mm.

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spur of hind tibiae widely spatuled. External margin ofelytrae posteriorly only slightly sinuate. Parameres in ventral view short, abruptly angulatedanteriorly, and stout (length/width almost 1), lobesrelatively long (ratio lobes length/parameres length:

0,43); aedeagus as in Fig. 8f.Female. Colouration as in male but antennae, labrumand maxillary palpi dark; fore margin of labrum dark;femurs and tibiae yellow-orange. Temples wide,slightly enlarged, maximum width of head on temples.Antennomere XI subquadrate.

Taxonomic remarks. The variety ruficornis, never col-lected after the description, represents only a phenotypewith the antennomeres completely red.

Distribution (Fig. 4). Mauritania. Ouadane (MNHN);Alzas. Embouchure (MNHN); Ile Tidra (MNHN);Mo-rocco. Sous, An Chab (Kocher, 1953; 1956); Arazane,Tifidelt (MNHN); Bou Tlidat (Kocher and Reymond,1954); Zguilma (ISR; Kocher, 1956); Algeria. Algeria(Kaszab, 1951; Dvorak, 1989); Bou Sada (MNHN);Hoggar, Tamouda (MNHN; Peyerimhoff, 1931); Biskra(Fairmaire, 1885; Bedel, 1895; CP; HNHM; MNHN);Ed Dor (HNHM); Haut Igharghar, Oued Tinbart (MNHN;Peyerimhoff, 1931); Haut Igharghar (MNHN); Amguid(MNHN; Peyerimhoff, 1931); Tifedest, AguelmameAraren (Peyerimhoff, 1931); Tunisia. Tunisia (Kaszab,

1951; Dvorak, 1989); Tozeur, road Nefta-Hazoua Km 20(CB); Khanget Oum-Ali (Bedel, 1895); Egypt.Egypt(Pic, 1923; Cros, 1939; Kasazb, 1951; Dvorak, 1989);Arabian desert, wadi E Maadi (Alfieri, 1976); Arabiandesert, wadi W Kosseir (Alfieri, 1976); Arabian desert,wadi Beida (Alfieri, 1976); Sinai (Chikatunov, 1999);Israel-Palestine. Dead Sea (Chikatunov, 1999); NorthernNegev (Chikatunov, 1999); Southern Negev (Chikatunov,1999); Arava Valley (Chikatunov, 1999).

Group hemprichi

Diaphorocera chrysoprasisFairmaire, 1863

Diaphorocera chrysoprasisFairmaire, 1863: 644.Diaphorocera kerimii Fairmaire, 1875: 530.

Type locality. Biskra (Fairmaire, 1863).D. kerimii: Gafsa (Fairmaire, 1875)

Type material. Fairmaire (1863) did not indicate where thefemale holotype is preserved, and we did not find it in the col-lections we examined.

The female holotype ofD. kerimii(MCSN) was examined.It has the following labels: Tunisia / Gafsa Aprile / AbdulKerim 1873 (white printed, except Gafsa Aprile, handwritten);4 (white, handwritten); Diaphorocera / kerimii / Fairm(white, handwritten); Typus (white, printed with red ink);Museo Civico / di Genova (white, printed). The holotype is in

good conditions even though both antennae are glued on thesame label of the specimen, except the first two antennomereswhich are still on head.

Diagnosis.Male. (Figs 6g; 7g; 8g; 9g; 10f, o; 11g, h, i).6-10 mm length. Green metallic, sometimes withgolden reflection on head and prothorax, rarely elytragreen-blue; antennae, mouthparts (except the externalside of mandibles), frons between antennal sockets,antero-lateral portion of head, anterior to eyes, and legs(except coxae) orange. Frontal calli with posterior margin on dorsal view,

reaching the posterior half of the margin of eye, slight-ly visible on lateral view; anterior portion evidentlyarcuate but not prolonged in the middle, externally notdifferentiated. Temples very elongate and parallel,slightly shorter than the longitudinal diameter of eye,the maximum width of head on eyes. Labrum about 2times as long as clypeus. Maxillary palpomere III sub-equal to II; distal end of III obtuse; IV longer than twiceits maximum width. Anterior side of antennomere IIIwith few setae, about as long as those on II; anteriormargin of IV, V and VII bilobate; posterior side of an-

tennomeres III-VIII with short setae; VIII-X largelytransverse and flattened, VIII distinctly wider than theothers, apically rounded, IX apically truncate, X half-moon like; width of X distinctly shorter than the lengthof XI; XI elongate. Pronotum narrow and elongate, anteriorly with twooblique and deep grooves. Fore tibiae dorsally flattenedwith a laminar expansion on the external side upwardcurved; external spur of hind tibiae narrowly spatuled.External margin of elytra posteriorly not evidentlysinuate. Lobes of parameres relatively short (ratio lobes

length/parameres length: 0,28); aedeagus apically stout,with sligthly elongate hooks.

Female. Colouration as in male, except mouthpartsslightly darker; frons between antennal sockets andclypeus basally metallic. Temples elongate and parallel;maximum width of head on eyes. Antennomere XIelongate.

Taxonomic remarks.D. kerimii was synonymised withD. chrysoprasisby Bedel (1895). After the examination


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of the holotype of the former taxon we confirmed thissynonymy with some doubts. In fact this specimen is afemale lacking both antennae; the general shape andseveral features agree with chrysoprasis, except thelarger size, and the pronotum wider and with sides not

converging basally. Several specimens from Gafsa - thetype locality ofD. kerimii- and adjacent zones wereexamined, all referable to typicalD. chrysoprasis. Some morphological differences withD. hemprichiare discussed below.

Fig. 5.Male head, dorsal view: (a) D. carinicollis, (b)D. johnsoni, (c)D. obscuritarsis, (d)D. promelaena, (e)D. peyerimhoffi, (f)D.sicardi, (g)D. chrysoprasis, (h)D. hemprichi. Scale bars = 1 mm.

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Distribution (Fig. 3).Mauritania. Khatt Atoui, Tamarat(MNHN); Algeria. Algeria (Kaszab, 1951; Dvorak,1989); Bou Sada (Pic, 1897; MNHN; HNHM); OuedChar (HNHM); Biskra (Fairmaire, 1863; HNHM;MNHN); Laghouat (HNHM); Touggourt (MNHN;

HNHM); An Sefra (HNHM; MNHN); Ghardaa(MNHN); Amguid, Tassili ouest (MNHN); Timuit(CB); Tunisia. Tunisia (Kaszab, 1951; Dvorak, 1989);S Kebili, Al Blidet (CBA); Od. Fekka (MHNG); Mak-nassy (Normand, 1949); Gafsa (Fairmaire, 1875;MCSN); Tozeur (Normand, 1949; MNHN); Tozeur, 10Km SW Tozeur on road P3 (CB); Tozeur, 25 Km fromTozeur on road P3 (CB); Nefta (CBA); El Golaa (Zaa-frane) (CS); 7 Km N Douz on road C206 (CB); Zarzis(MHNG; MNHN; HNHM); 30 Km W Nefta, Hazoua(CBA); Libya. Tripolitania, Al Aziziyah (CB; CR);Tripolitania, Suq Ai Vchamis (CK).

Diaphorocera hemprichiHeyden, 1863

D. hemprichi hemprichiHeyden, 1863: 127.

Type locality. D. hemprichi hemprichi: Aegypten(Heyden, 1863).

Type material. Types of the nominate subspecies were notexamined; they are probably preserved at the Zoological Museumof Berlin.

Diagnosis.Male. (Figs 3h; 4h 5h; 6h; 7g, d; 8h; 9l-n;11n). 6-12 mm length. Similar toD. chrysoprasis ex-cept for the following characters: temples shorter andslightly curved; antennomere VIII slightly wider thanIX and X; anterior portion of antennomeres VIII-IXapically narrowing; width of X slightly shorter than thelength of XI. Pronotum slightly wider than in male;foretibiae with a laminar expansion also on the internalside. Lobes of parameres raised at base and more di-verging externally.

Female. Colouration as in male, except mouthpartsslightly darker and clypeus basally metallic. Templesprogressively converging posteriad; maximum widthof head on eyes. Antennomere XI elongate. Pronotumslightly wider.

D. hemprichi sauditaKaszab, 1983: 179.

Type locality. D. hemprichi saudita: Saudi Arabia,Eastern Prov., Udhailiyah Camp (Kaszab, 1983).

Type material. Four paratypes of D. hemprichi saudita(HNHM) were examined. One male and one female have thesame following labels: Jaww Dukha / 7.V.1982 / no. 186 (white,printed); SAUDI ARABIA / Eastern Prov. / D. A. Pitcher(white, printed); Paratypus 1983 [andrespectively]/Dia-phorocera / hemprichissp. /saudita/ Kaszab (white with red

frame, handwritten, except Paratypus printed with red ink);Diaphorocera / hemprichi ssp . / saudita Kaszab / Dr. Z.Kaszab det., 1983 (white, handwritten, except the underlinedpart of the last line, printed); on an additional label hare gluedthe tegmen and penis, the last abdominal segment and the specu-lum gastrale. One third male has the same labels, but 28.VI.1982/ no. 192. The fourth male paratype has the following labels:Udhailiyah Camp / 29.IV.1982 / no. 171 (white, printed);SAUDI ARABIA / Eastern Prov. / D. A. Pitcher (white,printed); Paratypus 1983 /Diaphorocera / hemprichi ssp. /saudita / Kaszab (white with red frame, handwritten, exceptParatypus printed with red ink).

Diagnosis. Very similar to the nominate subspecies,being the differences, summarised by Kaszab (1983),very weak: (a) the head has a sparse puncturation in thenominate subspecies, but is almost smooth inD. hemp-richi saudita; (b) the pronotum is shorter and broaderin the former and more elongate and narrow in the lat-ter, moreover the female pronotum of the Arabian racehas punctures very sparse, uniform and fine.

Taxonomic remarks.D. hemprichiandD. chrysoprasisare very close and difficult to distinguish, except for the

antennal and head features listed in the diagnostic key.Kaszab (1951) emphasised the difference between thesespecies as concerns the inner side of male fore tibia.That ofD. hemprichiis expanded to form a lamina (Figs10g, p), but in the second species the inner margin issimple. Actually, in the Ain-Sfra (Algerian Sahara)population, living in syntopy withD. chrysoprasis, wefound a great variability of this character, and someindividuals have the inner side scarcely or not ex-panded.

Distribution (Fig. 11).D. hemprichi hemprichi.Morocco.

Draa Region, El Kantara (Pardo Alcaide, 1961); Seguiael Hamra, Hauga Ramala (Pardo Alcaide, 1961); OuedHasi Belibilia (ISR); Bou-Denib (Kocher, 1956); DraaValley, Agdz (CB); Gara Sbaa, Gara Bou Tlidat (ISR);Zguilma (Kocher, 1956); Rio de Oro, Smamit, UadKomba (Pardo Alcaide, 1961); Algeria. Bou Sada(MNHN); Oued Char (HNHM); Les Terres Blanches(CP); An Sefra (ISR; MNHN); Ghardaa (MNHN);Amguid (Peyerimhoff, 1931); Tunisia. Gabes (MVE);Egypt. Egypt (Heyden, 1863; Cros, 1939; Kaszab, 1951;HNHM); Libyan desert, Abu Rauwash (Alfieri, 1976);


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Libyan desert, Kirdasah (Alfieri, 1976; CB); Saqqarah(Pic, 1900); Masarah (MVE); Cairo, Mena, Hotel Gar-den (OUM); Arabian desert, wadi SE Cairo (Alfieri,1976); Israel-Palestine. Jordan Valley (Chikatunov,1999); Southern Coastal Plain (Chikatunov, 1999);

Judean Desert (Chikatunov, 1999); Northern Negev(Chikatunov, 1999); Gevulot (Kaszab, 1957); Urim(Kaszab, 1957).

D. hemprichi saudita. Iran. Abbassi, Bandar Abbas(Dvorak, 1996); 16 Km E Bandar Abbas (HNHM);Saudi Arabia. Saudi Arabia (Dvorak, 1989); Hofuf (Sch-neider, 1991); Eastern Province, Udhailiyah Camp(Kaszab, 1983); Jaww Dukha (Kaszab, 1983); Dibba(MNHN); Oman. Ras Dhabdhub (Schneider, 1991);United Arab Emirates. Al Jazirat Al Hamra Emirate,crossroad to Airport on road E11 (CB); Ras Al Khaimah

Emirate, 23 Km from Airport croosroad on road E18(Habab-Manama) (CB); Al An, Al Jahar-Al Saad on roadAl An-Abu Dhabi (CB); Abu Dhabi Emirate, Al An,Bateen Dunes (CB; CK); Abu Dhabi Emirate, Al An,3-4 Km E Al Selimat, on road Al An-Abu Dhabi (CB).

Diaphorocera promelaenaFairmaire, 1876

Diaphorocera promelaenaFairmaire, 1876: 49.

Type locality. Entre Bou-Sada et Biskra (Fairmaire,1876).

Type material. Two males and seven females paratypes(HNHM) were examined. All the paratypes have the same fol-lowing labels: Biskra / R. Oberthr / 1875 (white with thinblack frame, printed); coll. R. Oberthr / ex coll. Deyrolle(white, printed); Paratypus 1876 /Diaphorocera / promelaena/ Fairmaire (white with red frame, handwritten, except Paraty-pus printed with red ink). The holotype was not found. Threefemales lack some parts of the body: the last two tarsomeres ofhind left leg and the whole tarsus of hind right leg; the wholetarsus of hind left leg; the whole tarsus of hind right leg and

tibia and tarsus of hind left leg.

Diagnosis.Male. (Figs 3d; 4d; 5d; 6d; 7dm; 9a-c; 12c).6-10 mm length. Head, including the base of clypeusand mandibles, the remaining mouthparts black or dark,pronotum, abdomen, thoracic sternites, coxae, trochant-ers and antennomere I subopaque black; the rest of thebody blue-green metallic with antennomere II-XI, fe-murs, tibiae and tarsi (or partially dark) orange. Bodywith white setation, particularly long on templa, prono-tum and venter.

Head evidently transverse. Frontal calli bulged,evidently visible on lateral view, posterior margin, ondorsal view, reaching the posterior half of the marginof eye; anterior portion arcuate but not prolonged inthe middle, externally not differentiated. Temples

parallel and about as long as 2/3 of longitudinal di-ameter of eye; maximum width of head on eyes. La-brum about 1,5 times as long as than clypeus. Maxil-lary palpi III shorter than II, distal end obtuse; IVlonger than twice its maximum width; anterior side ofIII with few setae, shorter than those on II. Anteriormargin of antennomeres IV, V and VII not bilobate;posterior side of III-VIII with withish setae, particu-larly long on VI-VII; VII-IX very transverse vari-ously shaped, X triangular, shorter than IX; XI elon-gate, about as long as 2 times the width of X. Pronotum wide with fore half greatly curved, ante-riorly with two oblique and deep grooves. Fore tibiaemodified, with a laminar external expansion only onits distal half; external spur of hind tibiae narrowlyspatuled; claws short and curved. External margin ofelytra posteriorly slightly sinuate. Lobes of parameres elongate (ratio lobes length/parameres length: 0,38); phal lobase poster iorlyabruptly narrowed; aedeagus short and oblique at apex,hooks very short and differently sized; hooks paralleland oblique.

Female. Colouration and head size as in male. Anten-nomere XI elongate.

Distribution (Fig. 12). Morocco. Morocco (Dvorak,1989); Moyenne Moulouya, Tamdafelt (Kocher, 1954;1956; ISR); Moyenne Moulouya, Ouizret (Kocher,1954; 1956; ISR); Moyenne Moulouya, Amersid (Ko-cher, 1954; 1956; ISR); Oued Guir (MNHN); Taourirt(MNHN); Algeria. Algeria (Kaszab, 1951; Dvorak,1989); Bou Sada (Bedel, 1895); between Bou Sadaand Biskra (Fairmaire, 1876); Biskra (Bedel, 1895;

Baudi di Selve, 1878; CP; HNHM; MNHN); Laghouat(MNHN); An Sefra (MNHN); Ghardaa (MNHN);Colomb-Bchar (MNHN); Tunisia. Tunisia (Kaszab,1951; Dvorak, 1989); El Kef (Normand, 1938); Mak-nassy (Normand, 1949); Tamerza (CBA); Metlaoui(CB); Gabs (Normand, 1949; MHNG; MNHN;MVE); Zarzis (MNHN); Oglet El-Rechid (El Borma)(Bedel, 1895);Egypt. Sinai, Wadi Isla (Alfieri, 1976);Sinai (Chikatunov, 1999). The Alfieris record, prob-ably re-cited by Chikatunov (1999), need confirma-tion.


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Fig. 6. Male pronotum, dorsal view: (a)D. carinicollis, (b)D. johnsoni, (c)D. obscuritarsis, (d)D. promelaena, (e)D. peyerimhoffi, (f)D. sicardi, (g)D. chrysoprasis, (h)D. hemprichi. Scale bars = 1 mm.

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Diaphorocera sicardiBedel, 1917

Diaphorocera sicardiBedel, 1917: 364.

Type locality. Maroc oriental: Fritissa (Bedel, 1917).

Type material. The male holotype (MNHN), which was exam-ined, has the following labels: Fritissa/Maroc (white, handwrit-ten); type (white, printed); Diaphorocera / sicardi / Bedel /n.sp. (white, handwritten); MUSUM PARIS/1930/coll. Si-card (white, printed).

The holotype lacks the right middle leg, except the coax, stillpresent.

Diagnosis.Male. (Figs 3f; 4f; 5f; 6f; 7e, n; 9d-f; 10).9-15 mm length. Blue-green metallic with head, thoraxand abdomen darker, mandibles and frons betweenantennal sockets dark; clypeus, labrum, antero-lateralportion of head anterior to eyes, antennae and legs(except coxae and trochanters black and tarsi dark,posterior tibiae sometimes dark) orange; the extensionof black colouration on middle and hind legs varies insome specimens, in which is extended on tarsi and mostpart of tibiae. Labrum with abundant and long yellow-ish setae. Head transverse, with a slight depression posteriorto calli and on temples, consequently vertex appearsslightly bulged. Frontal calli clearly visible on lateralview; posterior margin on dorsal view reaching the

posterior half of the margin of eye; anterior portionevidently arcuate and prolonged in the middle withtwo distinct pointed protrusions, externally differenti-ated with two elevated keels, apically pointed and an-teriorly protruding; apexes of frontal keels slightly in-wardly directed (Fig. 8f). Temples slightly enlarged,about as wide as eyes. Labrum about 1,5 times as longas clypeus, with an longitudinal laminar appendix,forming an arch, extended from the apex posteriad.Maxillary palpomere III subequal to II; distal end ofIII externally elongate and pointed; IV about twice aslong as its maximum width and swollen after the mid-

dle. Antennomeres IV-X transverse; anterior portion ofIV bilobate, basal lobe pointed and curved; anteriorportion of VII with a long dorsal spine-like protrusion;VIII-IX with a fore chitinous line, X with a reticulationof chitinous lines and spots, XI with single chitinousspots at the base; VIII subfalcate, IX falcate, X widelysubtrapezoidal, XI very transverse. Pronotum elongate, subparallel sided, anteriorlyslightly angulated with two oblique and deep grooves.Fore tibiae on lateral view with an external short in-flated expansion, with a corresponding internal depres-

sion, on lateral view gibbose at base; external spur ofhind tibiae narrowly spatuled. External margin of elytraposteriorly not evidently sinuate. Lobes of parameres relatively short (ratio lobeslength/parameres length: 0,28); aedeagus apically stout,

with sligthly elongate hooks.

Female. Colouration as in male, except mouthpartsslightly darker; frons between antennal sockets andbased of clypeus metallic. Temples elongate and paral-lel; maximum width of head on eyes. Antennomere XIelongate.

Taxonomic remarks. One female, identified by Kaszab(HNHM), from Tozena, an undetected localit, probablyof Algeria, differs because of the head and pronotumshiny black, sides of pronotum parallel, antennae and

fore tibiae dark.

Distribution (Fig. 10). Morocco. Morocco (Kaszab,1951; Dvorak, 1989); Guercif (MNHN); Fekkous(Kocher, 1954; 1956; ISR); Fritissa (Bedel, 1917;Kocher, 1956); Algeria. Oum Ali (HNHM); Tunisia.Gafsa (MHNG);Libya. Tripolitania, Tasvadi-Mesari,Bin Ghashir (CK).

Diaphorocera peyerimhoffiKocher, 1954

Diaphorocera peyerimhoffiKocher, 1954: 284.

Type locality. Ouizret (Kocher, 1954).

Type material. Two males (Type and Cotype) and one female(Allotype) were examined (ISR); they have not an identificationlabel on the pin, which was in the box, just close to the specimens.The type has the following label: Ouizret/ Moy. Moulouya /(Kocher) 5.53 (white, handwritten, except Moy. Moulouyaprinted); (white, printed); TYPE (red, handwritten). Thecotype has the following label: Ouizret / (Moy. Moulouya) /Kocher 5/53 (white, handwritten, except (Moy. Moulouya),

Kocher and 53 printed); COTYPE (red, handwritten). Theallotype has the following label: Ouizret / (Moy. Moulouya) /Kocher 5/53 (white, handwritten, except (Moy. Moulouya),Kocher and 53 printed); (white, printed); ALLOTYPE(red, handwritten).

All the specimens were in good conditions.

Diagnosis.Male.(Figs 3e; 4e; 5e; 6e; 10). 13-17 mmlength. Similar toD. sicardi except for the followingcharacters: body larger in size; temples evidentlywider and enlarged; frontal calli with anterior portionevidently arcuate but not greatly prolonged in the mid-


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dle, with externally protrusion externally directed (Fig.8e); apex of maxillary palpomere III more pointed andIV lesser swollen; antennomere IV width basal lobeuncinate, VII bilobed, VIII-IX not falcate, X narrowand subfalcate, chitinous areas absent.

Female. Similar toD. sicardi,but body larger in size;frons with a red spot.

Taxonomic remarks. Diagnostic characters, additionalto the Kochers description (1954), were detected; theypermit to clarify the phylogenetic position of this speciesand to discriminate it fromD. sicardi. In addition to thefeatures used in the key to the species, the followingcharacters could be evidenced. Both species have the synapomorphic condition offrontal calli with a raised external keel. InD. peyerim-

Fig. 7.Male right fore tibia, dorsal view: (a)D. carinicollis, (b)D. johnsoni, (c)D. obscuritarsis, (d)D. promelaena, (e)D. sicardi, (f)D. chrysoprasis, (g)D. hemprichi; lateral view: (h)D. carinicollis, (i)D. johnsoni, (l)D. obscuritarsis, (m)D. promelaena, (n)D. si-

cardi, (o)D. chrysoprasis, (p)D. hemprichi. Scale bars = 1 mm.

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hoffi the fore margin of frontal callus is internallyprolonged with a low and short protrusion, and exter-nally by the apex of a raised keel outwardly bent (Fig.8e); on the contrary, inD. sicardithe internal protrusionis more extended and the apex of the external keel isforwardly direct (Fig. 8f). The anterior margin of anten-nomere V is truncate, while in D. sicardi is pointed.Temples ofD. peyerimhoffiare enlarged posteriad andslightly wider than the width of head on eye, while inD. sicardiare parallel and about as wide as the widthof head on eye.

Distribution (Fig. 10).Morocco. Tahiant (Kocher, 1956;MNHN); n Thala, Tizi Talzent (ISR); Ouizret (Ko-cher, 1954; 1956); Haut Ziz, Rich (Kocher, 1956; ISR);Mzizel (ISR); Haut Ziz, nZala (Kocher, 1956; ISR).

Biogeography

The entire tribe Cerocomini is primarily a South Palae-arctic group, with an extension in East Africa of a newgenus related to Diaphorocera (Turco and Bologna,

Fig. 8.Male genitalia: D. carinicollis(a) tegmen, ventral view, (b) lateral view, (c) aedeagus, lateral view; D. johnsoni(d) tegmen,ventral view, (e) lateral view, (f) aedeagus, lateral view;D. obscuritarsis(g) tegmen, ventral view, (h) lateral view, (i) aedeagus, lateralview. Scale bars = 1 mm.

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unpublished). The most genera are West Palaearctic,and only one species of Cerocoma, C. schreberi Fabri-cius, 1781 is widely distributed from the Iberian Penin-sula to W China. Rhampholyssa is distributed in thesteppe and deserts of the Don and Turan lowlands,Rhampholyssodes is endemic to the Arabian deserts,Anisarthrocera, in the present meaning (see Classifica-tion) includes one Iraqi-Iranian species, and two EastAfrican species belong to the new undescribed genus.

Diaphorocera, is an eremic element, widely distrib-uted from the Atlantic coasts of Western Sahara and

Mauritania, through the entire Sahara and the Arabianpeninsula, to the southern Palestinian area and to thedry regions of South Iran. In synthesis it is a typicalSaharo-Sindian element (sensu Vigna Taglianti et al.,2000). Its possible relationships with an east Africangenus (Turco and Bologna, unpublished), one speciesof which (Anisarthrocera semirufa (Fairmaire, 1882)is specialised to semi-desert habitat, suggest interestinghypotheses of a common eremic ancestor of these twogenera, split in the Sahara and East Africa, probablyduring the a more humid Pliocenic phase.

Fig. 9. Male genitalia:D. promelaena(a) tegmen, ventral view, (b) lateral view, (c) aedeagus, lateral view;D. sicardi(d) tegmen, ventralview, (e) lateral view, (f) aedeagus, lateral view;D. chrysoprasis(g) tegmen, ventral view, (h) lateral view, (i) aedeagus, lateral view;D.hemprichi(l) tegmen, ventral view, (m) lateral view, (n) aedeagus, lateral view. Scale bars = 1 mm.

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Fig. 10.Distribution ofD. chrysoprasis(circles),D. peyerimhoffi(stars) andD. sicardi(triangles).

Fig. 11.Distribution ofD. hemprichi (triangles),D. obscuritarsis (circles) andD. johnsoni (stars).


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The phylogenetic relationships among the speciesemerging from the cladistic study, (see Classification)prompt some biogeographical considerations. The mostspeciose area results the Western part of Sahara, whereare distributed 7 species on 8 (Figs 3-5). One species,D. hemprichi, has a wide Saharao-Sindian distribution,

from Atlantic coasts to southern Iran; other specieshave a wide trans-Saharan range, asD. chrysoprasis,D. obscuritarsis, and possibly alsoD. promelaena (seeCatalogue), being spread from Morocco to Egypt.The in-group analysis of two main phyletic lineagessuggests the occurrence of late vicariance events, whichsplit these groups in separate areas: (a) In the obscuri-tarsisgroup, one species -D. obscuritarsis -is widelydistributed, but the remaining two, namelyD. carinicol-lis and D. johnsoni, are respectively endemic to theopposite sides of the Sahara-Arabic desert. (b) In thehemprichigroup, the robust cladeD. sicardi-D. peyer-imhoffi, includes one species more spread from Mo-rocco to Tripolitania (D. sicardi) and the second en-demic to a narrow area of the Moulouya Valley, in eastMorocco.

Acknowledgements

We wish to thank the following naturalists for the permission tostudy the Meloidae preserved in their Institutions or private col-lections and for their help: Claude Girard, Lionel Casset, Jean-

Claude Ringenbach, Paris; Roberto Poggi, Genova; Mauro Bon,Venezia; Otto Merkl, Budapest; Josef Jelinek, Jiri Hajek, JanBatelka, Praha; Stanislav Krejcik, Unicov, Czech Republic;Ahmed El Hassan and Mohamed Mouna, Rabat; John Pinto,Portland, Oregon, USA. We also thank Chiara Settanni, Serena Carloni and Andrea DiGiulio (Roma) for their help in the field research. Waleed Hamza

of the Al An University kindly supported our research in theUnited Arab Emirates. This study was supported by grants from the University RomeTre, Department of Biology (ex 60% and Programmi diricerca scientifica di rilevante interesse nazionale, n. 99C5271884-007) and from the Ministero dellIstruzione, dellUniversit edella Ricerca (coordinator Marco A. Bologna, n. 2004057217).We wish also to acknowledge the financial support of the Euro-pean Commissions Research Infrastructure Action via theSYNTHESYS Project, thanks to which researches in someEuropean Museums was carried out by one of us (FT) (FR-TAF-44 and HU-TAF-288).

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Received: 1 August 2006

Accepted: 23 March 2007


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Head

11 Frons between antennal sockets: dark (0), light (1)12 Maximum length of labrum vs that of clypeus: 1.5

times X as long (0), twice as long (1)13 Maximum width on eyes vs that on temples: great-

er (0), subequal or narrower (1)14 Posterior margin of frontal calli in dorsal view:

reaching the posterior half of eye length (0), reach-ing the middle of eye length (1)

15 Length of maxillary palpomere IV vs its maximumwidth: twice as long (0), more than twice as long(1)

16 Length of maxillary palpomere III vs that of segmentII: shorter (0), subequal or longer (1)

17 Shape of antennomere XI: elongate (0), subquadrate(1)18 Colour of antennomere I: dark, at least at base (0),

yellow-orange (1)19 Anterior margin of antennomere IV: bilobate (0),

not bilobate (1)10 Anterior margin of antennomere VII: bilobate (0),

not bilobate (1)11 Frontal calli, in lateral view: slightly visibile (0),

evidently visibile (1)12 Long and light setae on the posterior portion of

antennomeres VI-VII: absent (0), present (1)

13 Shape of anterior part of frontal calli: simple (0),raised with two keels apically pointed (1)

14 Colour of the antero-lateral part of head, anterior toeyes: completely dark (0), at least partially yellow-

orange (1)15 Shape of distal end of maxillary palpomere III:

simple (0), externally pointed (1)16 Setae on labrum: normal (0), long and lanuginose

(1)17 Labrum with a longitudinal medial area: slightly

raised (0), distinctly raised, separated and arcuate(1)

Thorax

18 Colour of middle and hind trochanters: yellow-or-

ange (0), metallic or black (1)19 Fore tibiae: simple (0), variously modified (1)20 Shape of fore tibiae: sub-cylindrical (0), with a

basal external short inflated expansion (1), with alaminar external expansion extended to the wholelength (2), with a laminar external expansion onlyat apex (3)

21 Shape of hind tibiae external spur: narrowly spatu-late (0), widely spatulate (1)

Abdomen

22 Number of hooks on aedeagus: one (0), two (1)

23 Number of hooks on endophallus: one (0), two (1)24 Ratio of lobes of parameres length vs parameres

length: less than 0.3 (0), more than 0.3 (1)

Appendix 1.Data matrix for the 24 characters used in the phylogenetic analysis.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24OUTGROUP 1 0 1 1 0 1 0 1 0 1 1 1 0 1 0 0 0 0 0 0 1 0 0 1D. hemprichi 1 1 0 0 1 1 0 1 1 0 1 1 0 1 0 0 0 0 1 2 0 1 1 0D. chrysoprasis 1 1 0 0 1 1 0 1 1 0 1 1 0 1 0 0 0 0 1 2 0 1 1 0

D. promelaena 0 0 0 0 1 0 0 0 0 1 1 1 0 0 0 0 0 1 1 3 0 1 1 1D. sicardi 1 0 1 0 0 1 0 1 1 0 1 0 1 1 1 1 1 1 1 1 0 1 1 0D. peyerimhoffi 1 0 1 0 0 1 0 1 1 0 1 0 1 1 1 1 1 1 1 1 0 1 1 ?D. obscuritarsis 0 0 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 1D. johnsoni 0 0 0 1 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1D. carinicollis 0 0 1 1 0 0 1 0 0 1 0 0 0 0 0 0 0 1 0 0 1 1 1 1

Appendix 2.Characters unordered used in the phylogenetic analysis.

Appendix 3. Cladogram statistics for the phylogenetic analysis.

Consistency index (CI) = 0.7429Homoplasy index (HI) = 0.2571CI excluding uninformative characters = 0.7273HI excluding uninformative characters = 0.2727Retention index (RI) = 0.8200Rescaled consistency index (RC) = 0.6091


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